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This article appeared in a journal published by Elsevier. The attachedcopy is furnished to the author for internal non-commercial researchand education use, including for instruction at the authors institution
and sharing with colleagues.
Other uses, including reproduction and distribution, or selling orlicensing copies, or posting to personal, institutional or third party
websites are prohibited.
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General palaeontology, systematics and evolution (Invertebrate palaeontology)
Cryptohelops menaticus – a new genus and species of the tribeHelopini (Coleoptera: Tenebrionidae) from the Palaeocene ofMenat (France)
Cryptohelops menaticus – un nouveau genre et une nouvelle espèce dela tribu des Helopini (Coleoptera : Tenebrionidae) du Paléocène deMenat (France)
Maxim V. Nabozhenkoa,∗, Alexander G. Kirejtshukb,c
a Murmansk Marine Biological Institute of Kola Scientific Centre RAS, Institute of Arid Zones of Southern Scientific Centre RAS, SouthernFederal University, Chekhov street, 41, 344006 Rostov-on-Don, Russiab Zoological Institute RAS, Universitetskaya nab., 1, 199034 St. Petersburg, Russiac CNRS UMR 5202, Muséum national d’histoire naturelle, CP 50, Entomologie, 45, rue Buffon, 75005 Paris, France
a r t i c l e i n f o
Article history:Received 28 May 2013Accepted after revision 22 September 2013Available online 2 December 2013
Presented by Philippe Taquet
Keywords:PalaeoceneMenatTenebrionidaeHelopiniNew genus and speciesFrance
a b s t r a c t
A new genus and species, Cryptohelops menaticus gen. et sp. n., are described from thePalaeocene of Menat (France). The new genus belongs to the subtribe Helopina of the tribeHelopini (subfamily Tenebrioninae of Tenebrionidae) based on the coarse and dense punc-tation of the hypomera, shape of the prosternum and metepisterna, a similar appearanceto the native genera Helops and Stenohelops and the structure of the epipleura. The newgenus also resembles Raiboscelis and Entomogonus by having the protibiae excised alongthe base of the inner side. The new genus is the oldest representative of the tribe Helopini.Four species of Helops sensu lato previously described from the Oligocene (H. wetteravicusK. Heyden et L. Heyden, 1865) and Miocene (H. atticus Redtenbacher in Ungern, 1867;H. meissneri Heer, 1847 and H. molassicus Heer, 1883) lack important diagnostic charactersand should be regarded as members of the tribe Helopini with uncertain generic attribution.
Mots clés :PaléocèneMenatTenebrionidaeHelopiniNouveau genreNouvelle espèceFrance
r é s u m é
Un nouveau genre et une nouvelle espèce, Cryptohelops menaticus gen. et sp. n. sont décritsdans le Paléocène de Menat (France). Le nouveau genre appartient à la sous-tribu des Helop-ina de la tribu des Helopini (sousfamille Tenebrioninae des Tenebrionidae) sur la base d’uneponctuation dense et grossière de l’hypomera, de la taille des prosternum et metepisterna,apparence similaire à celle des genres natifs Helops et Stenohelops, et de la structure des epi-pleura. Le nouveau genre ressemble aussi à Raiboscelis et Entomogonus de par les prototibiaexcisés le long de la base du bord interne. Le nouveau genre est le plus ancien représentantde la tribu des Helopini. Quatre espèces de Helops sensu lato, décrites antérieurement, àl’Oligocène (H. wetteravicus K. Heyden et L. Heyden, 1865) et au Miocène (H. atticus Redtebbacher in Ungern, 1867 ; H. meissneri Heer, 1847 et H. molassicus Heer, 1883) ne comportent
66 M.V. Nabozhenko, A.G. Kirejtshuk / C. R. Palevol 13 (2014) 65–71
pas certains caractères diagnostiques importants et devraient être considérées comme desmembres de la tribu des Helopini à attribution générique incertaine.
The family Tenebrionidae is estimated at more than20 000 living species and is known from more than 100extinct species. The present paper describes a new genusand species of the tribe Helopini (subfamily Tenebrioni-nae sensu str.) from the Middle Palaeocene of Menat (seebelow). This work is the first of a series of publicationsdevoted to the “Menat” Coleoptera fauna.
Materials from this locality deposited in the MenatTown Museum and National Muséum d’Histoire Naturelle(MNHN) are considerably rich with insect remains but arevery difficult to study because of the fragility of the sub-strate and insect remains, and available specimens are forthe most part very incomplete (with missing appendagesand unclear segmentation). While the Coleoptera is thelargest group of all invertebrate remains obtained from thissite (nearly 70% of more than 2000 samples), most bee-tle remains can be identified only to the family level. Thematerials recently examined (including re-examination ofsome types deposited in NMHN) clearly demonstrate thelack of aquatic species (except two specimens of the fam-ily Gyrinidae (Piton, 1940; Nel, 1989) and one specimenof Dytiscidae) and absolute dominance of the family Cur-culionidae among the coleopteran families (more than halfof the beetle remains fall within this family). In spite oftheir great numbers, only a few rather numerous speciesindicate arboreal inhabitants. This is a unique locality withsuch a large proportion of weevils, although this familyis dominant in many Cainozoic deposits (Kirejtshuk andPonomarenko, 2012; Kirejtshuk et al., in preparation). Thenext most abundant coleopterous family is the Bupresti-dae (about 50 remains). This family is dominant only in theUpper Cretaceous outcrop in Zerkal’naya (Primorsky Kray,Russia: Ponomarenko, pers. comm.) and in the younger sitein Geisental (Middle Eocene) where Buprestidae occupy30–40% of the total number of beetles (Haupt, 1950, 1956;Hörnschemeyer et al., 1995). However, such high numbersseem to be based on particular circumstances because in allother Palaeocene environments the representation of thisfamily is not so significant (Kirejtshuk and Ponomarenko,2012). It is also significant that the most recent Bupresti-dae, probably like most fossil members of the family,are/were xylophagous as larvae. Also represented in thisfossil assemblage are members of the families Elateridae,Cerambycidae, possibly Chrysomelidae and Nitidulidae aswell as some others, all represented by species whichtoday are associated with forest habitats. The present studydescribes a tenebrionid from this deposit that possessessufficient diagnostic characters to determine its correctsystematic position.
2. Material and methods
The holotype of the species under considerationis deposited in the Menat Town Museum, Village of
Menat, Puy-de-Dôme. The specimen was studied usinga stereomicroscope Olympus SCX9 in MNHN, and also astereomicroscope Leica MZ 16.0 in the Zoological Institute(St.-Petersburg) with different modes of light and colorfilters. The specimen was also examined with a TescanVega LSU scanning electron microscope in MNHN to testthe characters not clearly visible in the optic stereomi-croscope (puncturation, margins of legs, pronotum, elytra)with the Low Vacuum Secondary Electron TESCAN Detector(LVSTD).
3. Geological setting and locality information
The Middle Palaeocene Menat fossil site (Menat Basin,Puy-de-Dôme, France) is a volcanic maar containing alake about 2–3 km in diameter, which at present containssedimentary rocks (spongo-diatomites) with remains ofdiverse aquatic and terrestrial flora and fauna (Nel, 1989,2008; Piton, 1940; Stroinski and Szwedo, 2012 etc.). Thislocality has been well known since the beginning of the20th century (Fritel, 1903; Laurent, 1912 etc.) and a gen-eral preliminary overview of its biota was published byPiton (1940). According to Wappler et al. (2009), the plantand insect diversity in the lacustrine outcrop of Menat iscomparatively high, confirming the results of Piton (1940),who was the first to describe the fossil flora and fauna. Thecomposition of faunal and floral remains makes it possibleto conclude that this lake was surrounded by a forest andthe palaeoenvironment was warm and humid. Followingthe pollen, paleomammalian stratigraphic, and radiomet-ric K/Ar analyses, the age of Menat was estimated as 59Ma (Kedves and Russel, 1982; Nel, 2008). However, thenew estimate based on macroflora postulated its age within60–61 Ma (Wappler et al., 2009).
Note. The beetle has the distinct characters of thesubfamily as well as those of the tribe Helopini; namelyan open mouthpart (the cardo and stypes are not closedby the mentum); the mentum widened anteriorly witha median elevation; eyes small and not strongly trans-versely extended on the underside of the head; the apicalmaxillary palpomere pelecoid; the antennae subfiliformand moderately thickened; the femora thickened; tibiaelong; protarsi widened; metatarsi long, with stronglyelongated tarsomeres 1 and 4; epipleura strongly widenedat base; striae deep; lateral margin of elytra widelysinuated; prohypomera with coarse large punctures and
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Fig. 1. Cryptohelops menaticus gen. et sp. n. from Palaeocene of Menat, holotype, body ventrally; A, B: photographs of holotype (stereomicroscope, differentcolor filters and direction of light), dry print; C: figured holotype; D: holotype, electronic microscopy. Scale bare is for all figures.Fig. 1. Cryptohelops menaticus gen.et sp.n. du Paléocène de Menat, holotype, vue ventrale du corps ; A, B : photographies de l’holotype (stéréomicroscopeavec différents filtres de couleur et directions de la lumière), pointe sèche ; C : dessin de l’holotype ; D : holotype vu au microscope électronique. Barred’échelle identique pour toutes les figures.
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Fig. 2. Cryptohelops menaticus gen. et sp. n. from Palaeocene of Menat, holotype, body ventrally; A: head (stereomicroscope), dry print; B: head andprothorax, electronic microscopy; C: meso- and metaventrite, electronic microscopy.Fig. 2. Cryptohelops menaticus gen. et sp. n. du Paléocène de Menat, holotype, vue ventrale du corps ; A : tête (stéréomicroscope), pointe sèche ; B : tête etprothorax, vus au microscope électronique ; C : méso et métaventrite, vues au microscope électronique.
membranes between abdominal segments 3-4 and 4-5 welldeveloped.
Cryptohelops gen. n. (Figs. 1 and 2)Type species: Cryptohelops menaticus sp. n.Etymology. The name of the new genus is formed
from the Greek “kryptos” (cryptus – secret; covert, veiled,
clandestine, stealthy, privy) and the generic name Helops;gender masculine.
Diagnosis. Body elongate-oval. Head somewhat hypog-nathous, with clearly visible gula and gular sutures atbase of epicranium. Laryngeal margin trisinuated, withprojected angles. Mentum transverse, cordiform, with dis-tinct longitudinal elevation in middle. Cardo and stypes
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Fig. 3. Cryptohelops menaticus gen. et sp. n. from Palaeocene of Menat,holotype, body dorsally (counterpart).Fig. 3. Cryptohelops menaicus gen. et sp. n. de Paléocène de Menat, holo-type, le dos du corps (homologue).
rounded, with distinct suture between them. Maxillaeelongate, with first palpomere elongate, second one trans-versely triangular and widened apically (weakly pelecoid),third one somewhat similar but larger. Outer marginof mandibles rounded, with subequal apical teeth. Eyesweakly transverse. Antennae moderately long, some-what longer than head and prothorax combined, weaklywidened from base to apex; scape well developed,antennomere 11 somewhat wider than antennomere 9;antennomere 10 shorter than each of both, weakly asym-metrical. Pronotum weakly cordiform. Prohypomera withvery coarse, round punctures; anterior part of prosternumwith less coarse and dense punctation. Procoxae round.Prosternal process narrow, widened apically, rimmed nearcoxal cavities. Femora thickened, tibia straight. Protibiadistinctly sinuate at base along inner edge. Outer edge ofmetatibiae with broad wavy crenulation. Epipleura wide inbasal quarter, uniformly narrowing posteriorly and reach-ing sutural angle of elytra. Striae clearly visible, deep.
Probable bionomy. Taking into consideration the bion-omy of the recent relatives and also the large proportionof forested specimens in the deposits of Menat, the typespecies of this new genus could also be associated withforest habitats.
Cryptohelops menaticus, sp. n.(Figs. 1–3)
Etymology. The name derives from the type locality“Menat”.
Locus typicus and stratum typicum. Menat Basin, Puy-de-Dôme, France. The Middle Palaeocene; approximately60–61 Ma. Type locality is situated around the small lakenear the village of Menat (46◦ 06′ N; 2◦ 54′ E). Map is givenin Stroinski and Szwedo (2012).
Type material. Holotype (male) deposited in MenatTown Museum, Village of Menat, Puy-de-Dôme: “MNT 05712A”.
The imprints of the holotype (part and counterpart)are well preserved. The part shows clearly visible ven-tral and partly dorsal surfaces of the body, antennae, headwith maxillae, mentum, mandibles (partly), eyes, gula withgular sutures, prosternum, prohypomera, all femora, oneprotibia with tarsus, both metatibiae with tarsi, punctationof prosternum, prohypomera, metepisterna, elytral striaewith punctures, epipleura. Also, the membrane betweenthe abdominal ventrites can also be seen (Fig. 1a, b, d).The counterpart demonstrates mostly outlines of the dorsalsclerites of the specimen and sculpture of their integument.
Description of holotype. Body oval. Head somewhatnarrower than prothorax, widest at level of eyes and tem-ples (genae destroyed). Eyes weakly transverse and weaklyconvex, antennae short, somewhat longer than prothoraxand head combined. Dorsal surface of head with com-paratively sparse and moderately large punctures, withintervals markedly greater (2–3 times) than a puncturediameter. Apical antennomere nearly twice as long asantennomeres 9 and 10 combined. Antennae slightly andgradually thickening apically. Temples behind eyes weaklyrounded, without visible grooves.
Pronotum cordiform, transverse, widest before middle;anterior edge very weakly and widely emarginate. Lat-eral edges weakly rounded in anterior half and widelysinuate in basal half. Surface of pronotum with compara-tively sparse and moderately large punctures, comparablewith those on head. Prosternum with moderately coarseand dense punctation (diameter of punctures subequal todistance between punctures). Prohypomera with coarseround dense foveae (punctures). Prosternal process notbordered apically.
Mesoventrite elongate, V-shaped depression not visi-ble. Metaventrite visibly transverse (twice as wide as long).Elytra in base clearly wider than pronotum, with welldeveloped humoral angles. Lateral sides of elytra broadlyrounded, sinuated near base. Elytral striae distinct, roundpunctures connected by narrow deep striae, interspacesbetween them with sparse and comparatively fine punctu-ation. Epipleura strongly widened basally and more narrowon toward apex, gradually reaching sutural angle of elytra.Abdominal ventrite 1 longer than following ones (espe-cially laterally), with acute intercoxal process. Anal ventritewidely regularly rounded at apex, distinctly bordered.
Legs long. Femora thickened; inner edge of profemorasinuate in apical part. Procoxal cavities bordered anteri-orly; mesocoxal cavities also bordered in posterior part.Intercoxal process between mesocoxae widely rounded.Protibiae straight, not widened apically, clearly excised inbasal quarter of inner edge. Metatibiae straight, with outeredge widely waved (crenulate), protarsomeres weakly
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distinctly widened, metatarsi long, about 3/4 as long asmetatibiae, metatarsomeres 1 and 4 longest.
Body length – 7.0, body width – 2.9 mm.
5. Comparison
The new genus belongs to the helopioid branch of thesubtribe Helopini, characterized by the absence of templegrooves, very coarse sculpture of deep fovea (punctures)of the prohypomera, completely rimmed marginal surfaceof the anal ventrite, and margination of the tibiae. Cryptoh-elops gen. nov. can be distinguished from the genera Helops,Stenohelops Reitter, 1922 and Probaticus Seidlitz, 1896 bythe body appearance and the structure of the epipleura,which is very thick at the base and very narrow apically. Thenew genus differs from all the above genera in the struc-ture of the asymmetric antennomere 11 (males of Helops,Stenohelops and Probaticus have very long antennae withelongate antennomere 11), protibiae excised along the baseof the inner edge and serrate metatibiae. The new genusadditionally differs from Stenohelops in the structure of itsprohypomera (Stenohelops has fine and sparse punctation)and has epipleura that reach the sutural angle.
The excised outline of the protibial base in Cryptoh-elops gen. nov. is similar to that in Raiboscelis Allard, 1876and Entomogonus Solier, 1848. But Raiboscelis and Ento-mogonus have clavate tibiae (Nabozhenko and Löbl, 2009;Nabozhenko and Tichy, 2011), while the new genus hascarving only in the basal quarter of its protibia. The some-what elongate and slightly asymmetric apical antennomereis similar to that in Raiboscelis, while many other genera ofHelopina have a long and modified antennomere 11. Thenew genus differs also from the two previous genera inthe obtuse small body. Iit also differs from Entomogonus inthe structure of the epipleura (Entomogonus has apicallywidened epipleura), the short antennae and the weaklytransverse eyes (Entomogonus has strongly transverse elon-gate eyes). From Raiboscelis, the new genus differs in thestructure of its humeral angles (Raiboscelis has very narrow,tooth-shaped shoulders) and general appearance, sinceRaiboscelis is more slender, elongate and subcylindrical.
The characters of the new genus, which are distinct fromall Helopini are the following: metatibiae widely undulat-ing along outer edge and maxillary palpomere 2 pelecoid.
The type species of this new genus cannot be comparedwith any of the fossil species described under “Helops”(see below) and differs from the recent species of HelopsFabricius, 1775. Thus, all the species of “Helops” can bepreliminarily regarded as questionable members of thisgenus. The new species differs from “Helops” wetteravi-cus K. Heyden et L. Heyden, 1865 in the conjoined striae;“H.” meissneri Heer, 1847 in the smaller body, longer andwider pronotum and apparently conjoined striae; “H.” atti-cus Redtenbacher in Unger, 1867 in the smaller body, andapparently conjoined striae. “Helops molassicus Heer, 1883remains unknown to the authors.
6. Discussion
After study of the mostly recent fauna of Tenebrion-idae Latreille, 1802 there are known 47 genera of the tribe
Helopini proposed for about 700 species in the Palaearc-tic Region, and also for about 80 described species fromAmerica (Nearctic and Neotropical Regions) and 5 speciesin other regions (Purchart and Nabozhenko, 2012). FossilHelopini from the Paleogene (Oligocene, Helops wetteravi-cus) and Neogene (Miocene, Helops meissneri, Helops atticus,Helops molassicus) were all assigned to the genus Helopssensu lato (Heer, 1847, 1883; Heyden and Heyden, 1865;Redtenbacher in Unger, 1867). Another Helops (originally“Helops sp.”) from the Baltic Amber was mentioned by Klebs(1910) and later by Larsson (1978). Kirejtshuk et al. (2008)published a review of all names proposed for fossil darklingbeetles and a more detailed review is given in the catalogueby Kirejtshuk and Ponomarenko (2012). The descriptionsof the compression fossil “Helops” spp. were based onquestionable features, such as the structure of the elytralstriae and occasionally the green color tinge of the elytralsurface (for example H. wetteravicus) and all these spec-imens should be regarded as species inquirendae. Helopsmeissneri is represented by only an elytral imprint withthe head and pronotum destroyed (Oeningen, Germany,Upper Miocene). Helops atticus was described based on theimprint of the right elytron (Kumi, Greece, Upper Miocene)and the author compared this species with Erionura gigan-tea (Kraatz, 1862) and Entomogonus saphyrinus (Allard,1876) (supposedly members of the subgenus EntomogonusSolier, 1848). Erionura Reitter, 1903 and Entomogonus rep-resent the genera close to Helops. Helops molassicus is alsoknown only by the imprint of an elytron (Laus, Greenland,Upper Miocene) and the description of Helops wetteravicusis based on the imprint of the right elytron (Salzhauzen,Germany, Upper Oligocene).
The subfamily Tenebrioninae is the largest group of thefamily but is still poorly represented in the fossil record. Fornow, only 10 of 28 tribes of Tenebrioninae are known fromfossils (Kirejtshuk and Ponomarenko, 2012; Kirejtshuket al., 2008: Alphitobiini Reitter, 1917; Amarygmini Gis-tel, 1856; Amphidorini LeConte, 1862; Bolitophagini Kirby,1837; Helopini Latreille, 1802, Opatrini Brullé, 1832;Palorini Matthews, 2003; Pedinini Eschscholtz, 1829; Tene-brionini Latreille, 1802; Toxicini Lacordaire, 1859 andUlomini Blanchard, 1845). Most members of these tribeswere described in the composition of the genera repre-sented in the Recent fauna, and only three generic nameswere proposed for the extinct species: Alphitopsis Kirejt-shuk, Nabozhenko et Nel, 2011 (Alphitobiini), ProteleratesWickham, 1914 (Amphidorini) и Eupachypterus Kirejtshuk,Nabozhenko et Nel, 2010 (Opatrini). Among the 35 fossiltenebrionins described in enough detail to be sure of theirattribution, the most numerous are Helopini (four speciesof Helops sensu lato), representatives of the opatrin lin-eage (three species of Opatrini from Gonocephalum Solier,1834, Ephalus LeConte, 1862 and Ulus Horn, 1870 and onespecies of Pedinini from Leichenum Dejean, 1834) and Boli-tophagini (three species of Bolithophagus Illiger, 1798 andRhipidandrus LeConte, 1862). The remaining tribes haveonly 1–2 species in the fossil record. The oldest speciesof the subfamily was described from Yixian, the Rubiconof the Jurassic and Cretaceous Yixian, Alphitopsis initialisKirejtshuk, Nabozhenko et Nel, 2011 (Alphitobiini). Takinginto consideration the records of other fossil tenebrionids
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(Alleculinae) in the Upper Jurassic (Medvedev, 1969), itcould be supposed that the subfamily Tenebrioninae had itsmain differentiations during the Upper Jurassic and LowerCretaceous. The oldest species of the tribe Helopini heredescribed demonstrates that in the Middle Palaeocene, thistribe was represented by forms that were rather similar tothe recent ones (and have similar specializations: widenedprotarsi, modified pro- and metatibiae, widened apical andpreapical palpomeres of maxilla). Such circumstances alsosupport the concept of Mesozoic differentiations of the sub-family Tenebrioninae, most of which seem to have beencompleted before the Cainozoic.
Acknowledgments
Valuable assistance in the preparation of this paperwas received from André Nel (Laboratoire de Entomolo-gie, MNHN). Sylvain Pont (Laboratoire de Minéralogie et deCosmochimie, MNHN) assisted the authors in their studyof the holotype by electronic microscopy. George PoinarJr. (Oregon State University) checked the English, makingit more fluent and transparent and also provided us withvery valuable comments. This work was supported by theprogrammes of “Research in Paris” of the City of Paris andvisiting professors of MNHN, and also from a grant of theRussian Foundation for Basic Research 12-04-00663-ɑ. Itwas performed in framework of program of the Presidiumof the Russian Academy of Sciences “Problems of the originof life and formation of the biosphere”. The authors greatlyappreciate valuable comments on an earlier version of themanuscript received from anonymous reviewers.
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